Middle Devonian-Early Carboniferous Conodont Faunas from the Khoshyeilagh Formation, Alborz Mountains, North Iran

Journal of Sciences, Islamic Republic of Iran 17(1): 53-65 (2006) http://jsciences.ut.ac.ir University of Tehran, ISSN 1016-1104

A.R. Ashouri*

Department of Geology, Faculty of Sciences, Ferdowsi University of Mashhad, Mashhad, Islamic Republic of Iran

Abstract Conodonts of Khoshyeilagh Formation, the most complete section of Devonian rocks in the Alborz Mountains, northern Iran, has been studied, and subspecies of the genera Icriodus, Bipennatus, Polygnathus, Pelekysgnathus, Ancyrodella, Palmatolepis, Bispathodus, Scaphignathus, Clydagnathus, Gnathodus, Scaliognathus and Dollymae have been obtained from the studied section. The faunas indicate an Eifelian to Tournaisian age. The falsiovalis, Late rhomboidea- Early marginifera, Latest marginifera-trachytera, Middle expansa-Middle praesulcata, bouckaerti and anchoralis-latus conodont zones have been identified in the formation. Six new species are described: Icriodus ariaei, Ic. alborzensis, Ic. khoshyeilaghensis, Polygnathus klapperianus, Pelekysgnathus housei and Pe. talenti.

Keywords: Conodonts; Devonian; Carboniferous; Iran; Alborz Mountains; Khoshyeilagh Formation

Later workers have proposed their own division of the Introduction formation. For example Brice et al. [14] suggested The existence of Middle and Upper Devonian rocks seven members and twelve “Beds” within the formation. in the eastern Alborz (Figs. 1 and 2) was firstly Comparison of previous studies and the current one is mentioned by Bozorgnia et al. [12] in an unpublished shown in Figure 3. In the present study the original report of the Iranian NationalArchive Oil Company; it was later ofsubdivisions SID suggested by Bozorgnia [13] are used, named Khoshyeilagh Formation and described by though the explained proposal of Brice et al. [14] is Bozorgnia [13]. It consists of 1345 m of carbonates perhaps a better representation of the sequence since it (mainly limestone), sandstone and shale with a rich is distinguishable simply in the field. fauna of brachiopods and less abundant tentaculitids and bryozoans. It is conformable with the Padeha Formation Stratigraphy below and the Mobarak Formation above. The Khoshyeilagh Formation was divided into six The ages of members 1 and 6 of the formation have members and seventeen “Beds” by Bozorgnia [13]. been challenged by previous researchers; the greatest

* E-mail: [email protected]

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Figure 1. Simplified geological map of part of the Central and Eastern Alborz Mountains.

controversy is concern with Member 1. Age suggested that most of Member 1 was Eifelian with, in assignments were given to the members on the basis of its upper part, some early Givetian limestone beds. brachiopods (Brice in [13]) as follows: Couvinian for Members 2 and 3 and most of Member 4 were believed Member 1, Givetian for Member 2, Frasnian for to be Givetian. The Givetian-Frasnian boundary occurs Member 3, Frasnian-Famennian for Member 4, in the upper part of Member 4. The Famennian begins Famennian for Member 5 and Late Famennian for with Member 6 and includes a third of Member 7 as Member 6. Bozorgnia [13] reported two algal intervals well. The upper two thirds of the Member 7 is Latest in the Khoshyeilagh Formation: Trochiliscus “Zone” Famennian (“Strunian”). from lower part of the section (Couvinian age according From conodont faunas and fish remains, [15] to brachiopods) and Umberella “Zone” an especially assigned an Early Devonian age to Beds 1-5 of [13] or distinctive interval in the Upper Devonian of Iran; it is Member 1 of [14]. They indicated a Givetian age for Frasnian-Famennian. Ahmadzadeh [2] studied conodont Member 3 and part of Member 4; Member 4 was and brachiopod faunas from the lower part of the assigned a Frasnian age, and a Famennian age was Khoshyeilagh FormationArchive (Member 1 up to the lower ofassigned toSID Members 5 to 7 of [14]. A question raised part of Member 3 of [13]). He defined Early Devonian by Hamdi and Janvier [15] about the lower contact of conodonts from the lower part of the Member 2; the Khoshyeilagh Formation. They believed a sequence conodonts from the upper part of the Member 2 were of 270 m overlain by the basal conglomerate of the thought to be not age-diagnostic. He discriminated two Khoshyeilagh Formation of [13], should be regarded as brachiopod biozones in the upper part of the formation, the lower part of the formation. Shahrabi made a similar one thought to be Early Frasnian, the other Middle to suggestion [5]. Brice et al. [14] considered the sequence Late Frasnian. underlying the Khoshyeilagh Formation to equate with Brice et al. [14] reported on the brachiopod and the Padeha Formation, whereas Hamdi & Janvier [15] bryozoan faunas but provided some information on did not accept this because of the lack of dolomite and corals and conodonts. On the basis of brachiopods, they gypsiferous intervals at its type locality.

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Figure 2. Index geological map showing the location of the studied area.

Previous Conodont Studies Biostratigraphy Ahmadzadeh [1] examined the Devonian- Reported conodonts here from “Bed” 10 to the top of Carboniferous boundary and Lower Carboniferous the formation are the first records of conodonts from strata in the Central Alborz, northern Iran, on the basis these horizons. Previous records cover up to “Bed” 9 [2] of brachiopods and conodonts. An Ordovician- and “Bed” 7 [15]. Devonian section S of ArchiveBojnurd in the eastern Alborz ofConodonts SID from Member 1 (sample S. 205) include was studied by Ahmadzadeh [3], again on the basis of species of Icriodus and Eognathodus. Eognathodus brachiopods and conodonts. His specimens have been bipennatus from “Bed” 2 of Member 1 probably reviewed by Weddige [16,17]. Ahmadzadeh et al. [4] indicates a Middle Devonian (pre-varcus Zone) age. gave a preliminary report on conodonts of the Despite 17 samples from Member 2, the faunas were Khoshyeilagh Formation from the Mighan section SW poor with only two samples (S.209 & S.210) producing of the type locality of the formation. Ashouri [6-11] species of Icriodus and Polygnathus, and another (S. studied Devonian-Carboniferous conodont faunas from 211) only conodont fragments. The predominant genera the Khoshyeilagh Formation and, in recent years much in Member 3 are Icriodus and Polygnathus. Of 19 unpublished research on conodonts from Iran has been samples from this member (S.226-S.244) the most undertaken by postgraduate students. important one (S. 241) has a rich fauna of Ancyrodella,

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Figure 3. Comparison of previous studies of the Khoshyeilagh Formation and the present study. The section and the lithological description is based on Bozorgnia [13].

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the species A. binodosa and A. pristina indicating the 4 displays distribution of conodont species and fasiovalis Zone (Lowermost asymmetricus Zone) of the subspecies in the present study. All studied samples in latest Middle Devonian-Early Frasnian. Thus the this research, obtained from the type section of the Middle-Upper Devonian boundary lies beneath of this Khoshyeilagh Formation, 105 km south of sample, possibly close to sample 240. The last sample Khoshyeilagh village (Fig. 2). from this member (S. 244) contains Icriodus iowaensis iowaensis indicating Early Famennian. Doubtless due to Family ICRIODONTIDAE MÜLLER & facies (mainly sandstones) only a few Icriodus and MÜLLER, 1957 Polygnathus specimens were obtained, not sufficient to Genus Icriodus BRANSON & MEHL, 1938 indicate precise horizons. Icriodus alborzensis n. sp. Bed 14 has no critical fauna; Beds 15 and 16 of [13] Plate 1, Figures 14-19 produced richer and a more diverse fauna including Icriodus, Polygnathus, Pelekysgnathus, Palmatolepis Derivation of name: From the Alborz Mountains, and, in its uppermost part, Clydagnathus. Sample 270 northern Iran. with Palmatolepis quadrantinodosa inflexa is the most Holotype: AFUM270, the specimen illustrated on important sample from this member. This species has a Plate 1, Figure 15 from sample S. 233. short range from the Upper rhomboidea to Lower Locus typicus and stratum typicum: The type marginifera Zone. Scaphignathus, mainly S. velifer locality of the Khoshyeilagh Formation (Fig. 2); (samples S. 276-S. 279), indicates the uppermost bioclastic limestone, 402 m above the base of the marginifera-trachytera zones for the upper part of formation (Fig. 3). Member 5. Diagnosis: A species of Icriodus with alternation of The last member of the formation, Member 6, lateral and medial rows of rounded, discrete denticles of includes a single bed (Bed 17 of [13]), containing similar size. The basal cavity is extended on the lower Polygnathus and Bispathodus. The uppermost strata of margin of the posterior part and shows a spur. The Member 6 produced species of Polygnathus, medial row of denticles is extended posterior to the Bispathodus, Gnathodus, Scaliognathus and Dollymae. lateral rows and there is a posterior cusp. The most important fauna of the uppermost Devonian Description: The middle row of denticles is was from sample S. 286. It produced Polygnathus extended posterior relative to the lateral rows of delicatus indicative of the old costatus Zone. Co- denticles by two to three denticles joined by a ridge; the occurrence of Bispathodus aculeatus aculeatus indicates most posterior denticle is the largest and forms a restriction to the Middle expana Zone. Despite lack of posterior cusp. evidence of facies changes, no conodonts were found in A single specimen (Pl. 1, Fig. 14) displays very the next three samples (S.287-S.289). These samples uniform and openly spaced denticles. One specimen (Pl. were from an interval with rich Late Devonian and 1, Fig. 15) has a more or less asymmetrical outline and a middle Tournaisian faunas in the underlying and flaring lower margin. Two specimens show closely overlying samples. The Devonian-Carboniferous spaced denticles; one of them (Pl. 1, Fig. 16) has a well- boundary thus lies somewhere in this interval. extended basal cavity in the posterior part and the other A Carboniferous fauna was obtained from S. 290. It (Pl. 1, Fig. 17) displays a distinct spur. Two specimens produced Dollymae bouckaerti, indicator of the show a wide basal cavity in the posterior part; one of bouckaerti Zone within the middle Tournaisian. A them (Pl. 1, Fig. 19) has a narrower spindle and narrow diverse conodont fauna from the last sample (S. 292), basal cavity. The other (Pl. 1, Fig. 18) is a broad form from the top of formation included species of with a large basal cavity, well developed spur and lacks Polygnathus, BispathodusArchive, Scaliognathus and ofa medial rowSID of denticles anteriorly. Gnathodus. The presence of Scaliognathus anchoralis Occurrence: Member 3 of the Khoshyeilagh europensis indicates the anchoralis-latus Zone the Formation; age Late Givetian. uppermost biozone of the Tournaisian. Material: 10 I elements.

Taxonomy Icriodus ariaei n. sp. Plate 1, Figure 12 Only pa elements have been described in the present study. The figured specimens herein are housed in the Derivation of name: In honor of Dr. Ali Asghar department of geology of Ferdowsi University of Ariaei, my first geological teacher who has spent about Mashhad, Mashhad, Iran with the prefix AFUM. Figure 50 years of his life on the geology of Iran.

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Sample No. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. S. TAXA 205 208 210 211 227 228 229 230 232 233 234 235 237 239 240 241 247 250 255 259 260 261 262 264 266 269 270 271 275 276 277 279 281 286 288 290 292 Bipennatus bipennatus Icriodus orri Icriodus obliquimarginatus Icriodus regularicrescens Icriodus arkonensis Polygnathus ling. ling. delta Polygnathus cf. perbonus Polygnathus alveolus Polygnathus xylus xylus Icriodus brevis Icriodus difficilis Icriodus alternatus alternatus Icriodus alborzensis Ancyrodella binodosa Ancyrodella pristina Polygnathus latus Icriodus subterminus Icriodus iowaensis iowaensis Icriodus costatus darbyensis Icriodus ariaei Polygnathus brevilaminus Pelekysgnathus housei Ancyrognathus sinelamina Icriodus expansus Polygnathus communis communis Polygnathus angustidiscus Pelekysgnathus talenti Polygnathus klapperianus Pelekysgnathus inclinatus Palmatolepis quadrantinodosa inflexa Polygnathus semicostatus Polygnathus perplexus Palmatolepis wolskaja Icriodus khoshyeilaghensis Clydagnathus ormistoni Scaphignathus velifer Clydagnathus cavusformis Clydagnathus darensis Polygnathus delicatus Archive of SID Bispathodus aculeatus aculeatus Polygnathus nodocostatus Pseudopolygnathus oxypageus Dollymae boukaerti Polygnathus bischoffi Gnathodus typicus Gnathodus pseudosemiglaber Scaliognathus anchoralis anchoralis Figure 4. Distribution of conodont species in the Khoshyeilagh Formation.

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Holotype: AFUM262, the specimen illustrated on Genus Pelekysgnathus THOMAS, 1949 Plate 1, Figure 12 from sample S. 260. Pelekysgnathus housei n. sp. Locus typicus and stratum typicum: The type Plate 3, Figure 1 locality of the Khoshyeilagh Formation (Fig. 2); Derivation of name: For Prof. Micheal House, my bioclastic limestone, 910 m above the base of the first conodont teacher who continually encouraged me formation (Fig. 3). and contributed greatly to knowledge of the Devonian. Diagnosis: A species of Icriodus characterized by an Holotype: AFUM797, the specimen illustrated on extremely large rounded basal cavity, tapering gradually Plate 3, Figure 1 from sample S. 260. anteriorly. The medial row of denticles is extended as a Locus typicus and stratum typicum: The type ridge posterior to the lateral denticles. Apart from the locality of the Khoshyeilagh Formation (Fig. 2); denticles in the three main rows, two further denticles bioclastic limestone, 910 m above the base of the are present aligned with the posterior denticles. formation (Fig. 3). Description: The middle row of denticles extends as Diagnosis: A species of Pelekysgnathus a ridge with a scalloped crest posteriorly continuing to characterized by a relatively circular outline in its the lower margin. The most posterior lateral row of posterior two-thirds, rather sharply converging in the denticles is aligned with two further denticles positioned anterior third, and with a triangular outline denticulation on one side. The denticles of the medial row are offset consist of a relatively strong, short and reclined cusp posteriorly relative to the lateral rows of denticles in a and apparently six denticles, irregular in form and size. closely spaced pattern. The unit is widest at the level of A thin ridge, visible on the left side of the cusp and just the most posterior lateral denticle. anterior of it, extends laterally downward nearly to one- Occurrence: Member 5 of the Khoshyeilagh third of the lower margin. Formation. According to the associated conodont fauna, Description: The species differs from Pe. furnishi by its age is Upper Devonian, Famennian, crepida Zone. having a more extended basal cavity and a small thin Material: 6 I elements ridge from one side; it lacks distinct transverse ridges intersecting the main row. Pe. glenisteri has two well- Icriodus khoshyeilaghensis n. sp. developed denticulated accessory processes in the Plate 1, Figure 13 posterior end of the main row and intersected denticles Derivation of name: Khoshyeilagh is the name of a in the main row. village in the Alborz Mountains, northern Iran, from Occurrence: Member 5 of the Khoshyeilagh which the formation takes its name. Formation. Based on the associated fauna, the species Holotype: AFUM259, the specimen illustrated on can be referred to the Late rhomboidea to the Early Plate 1, Figure 13 from sample S. 271. marginifera zones. Locus typicus and stratum typicum: The type Material: 8 I elements locality of the Khoshyeilagh Formation (Fig. 2); limestone, 1050 m above the base of the formation (Fig. 3). Pelekysgnathus talenti n. sp. Diagnosis: A species of Icriodus with a well Plate 3, Figure 2 developed posterior cusp and two lateral ridges of denticles without an extended medial row of denticles. Derivation of name: In honor of Prof. John Talent The basal cavity is moderately large. who has generously helped people around the world Description: The species does not show a discrete with knowledge of the Devonian. medial row of denticles. Its first few left-side denticles Holotype: AFUM798, the specimen illustrated on converge anteriorly andArchive form a “pseudo medial” row. ofPlate 3, Figure SID 2 from sample S. 262. The anteriormost three denticles of the lateral rows are Locus typicus and stratum typicum: The type fused with those of the medial row appear as crossed locality of the Khoshyeilagh Formation (Fig. 2); ridges. bioclastic limestone, 940.5 m above the base of the Occurrence: The species comes from the upper part formation (Fig. 3). of Member 5 of the Khoshyeilagh Formation. On the Diagnosis: A species of Pelekysgnathus with a basal basis of stratigraphical position, its age may be as young cavity of relatively quadrangular outline in its posterior as the crepida Zone. two-thirds, becoming much narrower in the anterior Material: 7 I elements third. The main row of denticles comprises a strong reclined cusp and only two strong denticles. Two thin

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ridges just anterior to the cusp extend laterally nearly Icriodus brevis STAUFFER, 1940 half way to the lower margin. Fig. 1. Upper view of AFUM103. S. 229. Description: This species is distinguished from Pe. Icriodus obliquimarginatus BISCHOFF & furnishi by having a strong cusp and only two strong ZIEGLER, 1957 denticles in the main row with no intersecting transverse Fig. 2. Upper view of AFUM111. S. 210. ridges. Pe. glenisteri has denticulated accessory Icriodus cf. regularicrescens BULTYNCK, 1970 processes and six or more denticles in the main row. Fig. 3. Upper view of AFUM113. S. 210. The species differs from Pe. housei in its outline and the Icriodus expansus BRANSON & MEHL, 1934 number of denticles in the main row. Fig. 4. Upper view of AFUM136. S. 261. Occurrence: Upper part of Member 5 of the Icriodus costatus darbyensis KLAPPER, 1958 Khoshyeilagh Formation. On the basis of the Fig. 5. Upper view of AFUM126. S. 259. stratigraphical position an early Famennian age is Icriodus difficilis ZIEGLER & KLAPPER, 1976 assigned. Fig. 6. Upper view of AFUM140. S. 235. Material: 8 I elements Icriodus orri KLAPPER & BARRICK, 1983 Fig. 7. Upper view of AFUM156. S. 210. Family POLYGNATHIDAE BASSLER, 1925 Icriodus arkonensis STAUFFER, 1938 Fig. 8. Upper view of AFUM166. S. 210. Genus Polygnathus HINDE, 1879 Polygnathus klapperianus n. sp. Icriodus cf. iowaensis iowaensis YOUNGQUIST & Plate 2, Figure 7 PETERSON, 1947 Fig. 9. Upper view of AFUM214. S. 255. Derivation of name: In honour of Prof. Gilbert Icriodus subterminus YOUNGQUIST, 1947 Klapper who has contributed importantly on Devonian Fig. 10. Upper view of AFUM237. S. 264. conodonts. Icriodus cf. raymondi SANDBERG & ZIEGLER, Holotype: AFUM316, the specimen illustrated on 1979 Plate 2, Figure 7 from sample S. 266. Fig. 11. Upper view of AFUM266. S. 266. Locus typicus and stratum typicum: The type Icriodus ariaei n. sp. locality of the Khoshyeilagh Formation (Fig. 2); Fig. 12. Upper view of holotype, AFUM262. S. 260. bioclastic limestone, 1001 m above the base of the Icriodus khoshyeilaghensis n. sp. formation (Fig. 3). Fig. 13. Upper view of holotype, AFUM259. S. 271. Diagnosis: A small species of Polygnathus with a Icriodus alborzensis n. sp. large depression occupying the anterior aboral surface. Fig. 14. Upper view of AFUM267. S. 234. Platform is rounded and flattened. Free carina distinctly Fig. 15. Upper view of holotype, AFUM270. S. 233. developed in the posterior part of the platform. Fig. 16. Upper view of AFUM271. S. 230. Description: The specimen is fragmentary, missing Fig. 17. Upper view of AFUM274. S. 233. small parts of the anterior end of the free blade. The Fig. 18. Upper view of AFUM267. S. 237. platform is rounded and flattened, with upturned outer Fig. 19. Upper view of AFUM275. S. 238. margin in the middle part. The ornament is shagreen- Icriodus alternatus alternatus BRANSON & MEHL, like on the outer margin. The carina bears strong 1934 denticles developed distinctly in the posterior part. The Fig. 20. Upper view of AFUM184. S. 229. aboral depression bears concentric striae and a strong Pelekysgnathus inclinatus THOMAS, 1949 keel gradually disappearing about the centre of the Fig. 21. Upper view of AFUM514. S. 269. depression. Large pit present at the conjunction of the Bipennatus bipennatus (BISCHOFF & ZIEGLER, platform anterior marginArchive with the keel. of1957) SID Occurrence: Upper part of Member 5 of the Fig. 22. Upper view of AFUM508. S. 205. formation; not older than marginifera Zone. Fig. 23. Upper view of AFUM504. S. 205. Material: 6 Pa elements Fig. 24. Upper view of AFUM505. S. 205. Fig. 25. Upper view of AFUM506. S. 208. EXPLANATION OF PLATES Fig. 26. Upper view of AFUM511. S. 208. Plate 1 Plate 2 Figures 1-3, 8, 15-18: ×100; Figures 4, 6, 7, 9, 10, 12, 14, 20, 21, 24, 26: ×80; Figures 5: ×50; Figures 11, Figures 1, 2, 13, 22: ×60; Figures 3, 7, 15: ×100; 13, 19, 22, 23, 25: ×60. Figures 4, 5, 9, 11, 14, 24, 25: ×50; Figures 6, 12, 16,

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18, 21: ×80; Figures 8, 10, 17, 19, 20, 23, 26: ×40. Fig. 22. Upper view of AFUM730. S. 279. Scaphignathus velifer HELMS, 1959 Polygnathus angustidiscus YOUNGQUIST, 1945 Fig. 23. Upper view of AFUM740. S. 276. Fig. 1. Upper-lateral view of AFUM300. S. 262. Fig. 24. Upper, lower views of. AFUM743. S. 279. Polygnathus linguiformis alveolus WEDDIGE, 1977 Bispathodus aculatus aculatus ZIEGLER, Fig. 2. Upper-lateral view of AFUM302. S. 210. SANDBERG & AUSTIN, 1974 Polygnathus weddigei (CLAUSEN, LEUTERITZ & Fig. 25. Upper view of AFUM688. S. 286. ZIEGLER, 1979) Fig. 26. Upper view of AFUM679. S. 286. Fig. 3. Upper-lateral view of AFUM303. S. 210. Polygnathus linguiformis linguiformis gamma Plate 3 morphotype KLAPPER, 1987 Fig. 4. Side view of AFUM304. S. 210. Figures 1, 10, 16: ×40; Figures 2, 11, 13, 17: ×80; Polygnathus cf. perbonus (PHILIP, 1966) Figures 3, 4, 6-9, 12, 14: ×60; Figure 5: ×50; Figure 15: Fig. 5. Side view of AFUM308. S. 210. ×100. Polygnathus xylus xylus STAUFFER, 1938 Fig. 6. Oblique upper view of AFUM315. S. 234. Pelekysgnathus housei n.sp. Polygnathus klapperianus n. sp. Figs. 1. a, b. Upper and lateral views of holotype, Fig. 7. Upper, side, oblique lower views of holotype, AFUM797. S. 260. AFUM316. S. 266. Pelekysgnathus talenti n.sp. Polygnathus brevilaminus BRANSON & MEHL, Figs. 2a, b. Upper and lateral views of holotype, 1934 AFUM798. S. 262. Fig. 8. Side view of AFUM326. S. 260. Ancyrognathus sinelamina (BRANSON & MEHL, Polygnathus alatus HUDDLE, 1934 1934) Fig. 9. Side view of AFUM337. S. 247. Fig. 3. Upper view of AFUM536. S. 260. Polygnathus semicostatus BRANSON & MEHL, 1934 Ancyrodella binodosa UYENO, 1967 Fig. 10. Side view of AFUM378. S. 271. Fig. 4. Side view of AFUM516. S. 241. Polygnathus cf. nodocostatus BRANSON & MEHL, Fig. 5. Upper view of AFUM520. S. 241. 1934 Fig. 6. Upper view of AFUM517. S. 241. Fig. 11. Upper view of AFUM393. S. 288. Fig. 7. Upper view of AFUM515. S. 241. Polygnathus bischoffi RHODES, AUSTIN & Fig. 8. Upper view of AFUM521. S. 241. DRUCE, 1969 Ancyrodella pristina KHALYMBADZHA & Fig. 12. Upper-lateral view of AFUM447. S. 292. CHERNYSHEVA, 1970 Polygnathus perplexus THOMAS, 1949 Fig. 9. Upper view of AFUM540. S. 241. Fig. 13. Side view of AFUM427. S. 271. Palmatolepis cf. wolskajae OVNATANOVA, 1969 Polygnathus communis communis BRANSON & Fig. 10. Upper view of AFUM630. S. 271. MEHL, 1934 Palmatolepis quadrantinodosa inflexa MÜLLER, Fig. 14. Side view of AFUM449. S. 269. 1959 Fig. 15. Side view of AFUM456. S. 291. Fig. 11. Upper view of AFUM552. S. 270. Fig. 16. Lower view of AFUM458. S. 229. Pseudopolygnathus oxypageus LANE, SANDBERG Polygnathus delicatus ULRICH & BASSLER, 1926 & ZIEGLER, 1980 Fig. 17. Upper view of AFUM465. S. 286. Fig. 12. Upper view of AFUM722. S. 290. Polygnathus sp. A Dollymae bouckaerti GROESSENS, 1977 Fig. 18. Upper view ofArchive AFUM491. S. 284. ofFig. 13. SIDUpper view of AFUM768. S. 290. Polygnathus sp. B Gnathodus pseudosemiglaber THOMAS & Fig. 19. Upper view of AFUM497. S. 265. FELLOW, 1970 Clydagnathus cavusformis RHODES, AUSTIN & Fig. 14. Upper view of AFUM767. S. 292. DRUCE, 1969 Fig. 15. Upper view of AFUM766. S. 292. Fig. 20. Upper view of AFUM723. S. 279. Fig. 16. Upper view of AFUM761. S. 292. Clydagnathus darensis RHODES, AUSTIN & Scaliognathus anchoralis europensis LANE & DRUCE, 1969 ZIEGLER, 1983 Fig. 21. Upper view of AFUM724. S. 281. Fig. 17. Upper, side and lower views of AFUM784. Clydagnathus ormistoni BEINERT, KLAPPER, S. 292. SANDBERG & ZIEGLER, 1971

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Acknowledgments Faunas from Iran. Ph.D. Thesis, University of Hull, 351 pp. (1990). The study was undertaken with the support of the 7. Ashouri A.R. The stratigraphical position of Member 1 Ferdowsi University of Mashhad, Iran. Late Prof. M.R. and Member 6 of the Khoshyeilagh Formation based on House (my mentor at the University of Southampton, conodont fauna and introducing three conodont zones UK), Dr. A. Ruttner (Vienna), Prof. P. Bultynck from Member 6. Geosciences 4: 64-71 (in Farsi) (1994). 8. Ashouri A.R. Middle-Devonian-Early Carboniferous (Belgium), Dr. K. Weddige (Germany), Prof. G. conodont fauna from the Khoshyeilagh Formation, Klapper (University of Iowa, USA) and Dr. C. Sandberg Alborz Mountains, north Iran. 15th International (Geological Survey, Colorado, USA) gave Senkenberg Conference, IGCP 421 & SDS Meeting, encouragement and help; Prof. J.A. Talent (Macquarie Frankfurt, 6 (2001). University, Australia) read the manuscript carefully; A. 9. Ashouri A.R. Late Devonian and Middle-Late Devonian Mousavi (Geological Survey, Tehran, Iran), A. conodonts from eastern and northern Iran. Revista Monazzami (Geological Survey, Mashhad, Iran) and E. Espaňola Micropaleontologia, 3: 355-365 (2004). 10. Ashouri A.R. Early Carboniferous (Tournaisian) Mohammadzadeh (Energy Ministry, Mashhad, Iran) conodont faunas from eastern and northern Iran. New were helpful during the research. This is a contribution Paleontology and Stratigraphy (in press a). to IGCP 421: North Gondwana mid-Palaeozoic bioevent 11. Ashouri A.R. Icriodus and Polygnathus (conodonts) from /biogeography patterns in relation to crustal dynamics. the Late Devonian of eastern Iran, and Middle-Late Devonian of northern Iran (in press b). References 12. Bozorgnia H., Bayat A., Memear A., and Mohajer M. Unpublished report. National Iran Oil Company (1970). 1. Ahmadzadeh H.M. Stratigraphische und Palaontologische 13. Bozorgnia F. Paleozoic foraminiferal biostratigraphy of Untersuchungen im Unterkarbon des zentral Elbors Central and East Alborz Mountains, Iran. National Iran (Iran). Clausthal Geol. Abh., 7, 104 pp. (1971). Oil Company, 4, 185 pp. (1973). 2. Ahmadzadeh H.M. Stratigraphie und Fauna im Devon des 14. Brice D., Jenny J., Stamfli G., and Bigey F. Le Devonian östichen Elburs Iran. Clausthal Geol. Abh., 23, 114 pp. de 1 Elbourz oriental: Stratigraphie, paléontologie (1975). (brachiopodes et bryozoaires), paléogeographie. Riv. Ital. 3. Ahmadzadeh H.M. Brachiopods and conodonts of south Paleont., 84: 1-56, Milano (1978). of Bojnurd and their stratigraphical results. Science 15. Hamdi B. and Janvier P. Some conodont and fish remains Faculty, Tehran University, 45: 12-24 (in Farsi) (1983). from Lower Devonian (lower part of the Khoshyeilagh 4. Ahmadzadeh H.M., Yazdi M., and Karimi L. Preliminary Formation) of north east Shahrud, Iran. Geol. Surv. Iran, analysis of Late Devonian conodonts from Khoshyeilagh 49: 195-210 (1981). Formation (Mighan section), eastern Alborz. Geological 16. Weddige K. New stratigraphic data on Devonian and Society of Australia Abstracts, 61: 43 (2000). Carboniferous formations from the Binalud and Ozback- 5. Alavi-Naeini M. Paleozoic Stratigraphy of Iran. Treatise Kuh Mountains N/E Iran. Ibid., 51: 105-116 (1983). on the Geology of Iran, Geol. Surv. Iran (in Farsi), 269 17. Weddige K. Zur Stratigraphie and Palaeogeographie des pp. (1993). Devons und Carbons von N/E Iran. Senck. Leth., 52: 179- 6. Ashouri A.R. Devonian and Carboniferous Conodont 223 (1984).

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